Editing Conifer cone

{{short description|Reproductive organ on conifers}}
{{for|cones in Cycadales|Cycad}}
[[File:Pinus coulteri MHNT Cone.jpg|thumb|A mature female [[Pinus coulteri|big-cone pine]] (''Pinus coulteri'') cone, the heaviest pine cone]]
[[File:Young spruce cone (2489694695).jpg|thumb|A young female cone on a [[Picea abies|Norway spruce]] (''Picea abies'')]]
[[File:Pinus cembra - male cones RHu 2023 02.JPG|thumb|Immature male cones of [[Pinus cembra|Swiss pine]] (''Pinus cembra'')]]

A '''conifer cone''', or in formal [[botany|botanical]] usage a '''[[strobilus]]''', {{plural form}}: '''strobili''', is a seed-bearing organ on [[gymnosperm]] plants, especially in [[conifer]]s and [[cycad]]s. They are usually woody and variously conic, cylindrical, ovoid, to globular, and have scales and [[bract]]s arranged around a central axis, but can be fleshy and [[Berry (botany)|berry]]-like. The cone of [[Pinophyta]] (conifer clade) contains the [[plant sexuality|reproductive]] structures. The woody cone is the female cone, which produces [[seeds]]. The male cone, which produces [[pollen]], is usually [[Ephemerality|ephemeral]] and much less conspicuous even at full maturity. The name "cone" derives from Greek ''konos'' (pine cone), which also gave name to the geometric [[cone]]. The individual plates of a cone are known as ''scales''. In conifers where the cone develops over more than one year (such as [[pine]]s), the first year's growth of a seed scale on the cone, showing up as a protuberance at the end of the two-year-old scale, is called an ''umbo'', while the second year's growth is called the ''apophysis''.<ref name="Rushforth">{{cite book | last=Rushforth | first=Keith | title=Conifers | publisher=Christopher Helm Publishers | publication-place=London | date=1987-01-01 | isbn=0-7470-2801-X | pages=221–223}}</ref>

The male cone ('''microstrobilus''' or '''pollen cone''') is structurally similar across all conifers, differing only in small ways (mostly in scale arrangement) from species to species. Extending out from a central axis are [[microsporophyll]]s (modified leaves). Under each microsporophyll is one or several [[sporangium|microsporangia]] ([[pollen]] sacs).

The female cone ('''megastrobilus''', '''seed cone''', or '''ovulate cone''') contains [[ovule]]s which when fertilized by pollen become seeds. The female cone structure varies more markedly between the different conifer families and is often crucial for the identification of many species of conifers.

==Female cone of the conifer families==

===Pinaceae===
<gallery>
File:Pinus sylvestris female strobilus and cone en.svg|Anatomy of a [[Pinus sylvestris|Scots pine]] (''Pinus sylvestris'') female strobilus.
File:Picea Pungens Young Cones.jpg|Young cones of a [[Picea pungens|blue spruce]] (''Picea pungens'')
File:Young Pine Cone Center.jpg|Cross-section of a young pine cone
</gallery>
The members of the [[pinaceae|pine family]] ([[pine]]s, [[spruce]]s, [[fir]]s, [[Cedrus|cedar]]s, [[larch]]es, etc.) have cones that are imbricate (that is, with scales overlapping each other like fish scales). These cones, especially the woody female cones, are considered the "archetypal" tree cones. The female cone has two types of scale: [[bract]] scale and seed scale (or ovuliferous scale), one [[subtended]] by each bract scale, derived from a highly modified [[branchlet]]. On the upper-side base of each seed scale are two ovules that develop into seeds after fertilization by pollen grains. The bract scales develop first and are conspicuous at the time of pollination; the seed scales develop later to enclose and protect the seeds, with the bract scales often not growing further. The scales open temporarily to receive pollen, then close during fertilization and maturation, and re-open at maturity to allow the seed to escape. Maturation takes 6–8 months from pollination in most Pinaceae genera, but 12 months in cedars and 18–24 months (rarely more) in most pines. The cones open either by the seed scales flexing back when they dry out, or (in firs, cedars and [[Pseudolarix|golden larch]]) by the cones disintegrating with the seed scales falling off. The cones are conic, [[cylinder (geometry)|cylindrical]] or [[ovoid]] (egg-shaped), and small to very large, from 2–60&nbsp;cm long and 1–20&nbsp;cm broad.

After ripening, the opening of non-[[serotinous]] pine cones is associated with their moisture content—cones are open when dry and closed when wet.<ref>Dawson, Colin; Vincent, Julian F. V.; Rocca, Anne-Marie. 1997. How pine cones open. Nature 390: 668.</ref> This assures that the small, windborne seeds will be dispersed during relatively dry weather, and thus the distance traveled from the parent tree will be enhanced. A pine cone will go through many cycles of opening and closing during its life span, even after seed dispersal is complete.<ref>{{cite web |url=http://www.newton.dep.anl.gov/askasci/bot00/bot00763.htm |title=Ask a Scientist: Pine Cone Wet and Dry |publisher=Newton.dep.anl.gov |access-date=2013-08-30 |archive-url=https://web.archive.org/web/20150226224440/http://newton.dep.anl.gov/askasci/bot00/bot00763.htm |archive-date=2015-02-26 |url-status=dead }}</ref> This process occurs with older cones while attached to branches and even after the older cones have fallen to the [[forest floor]].

The condition of fallen pine cones is a crude indication of the forest floor's moisture content, which is an important indication of [[wildfire]] risk. Closed cones indicate damp conditions while open cones indicate the forest floor is dry. As a result of this, pine cones have often been used by people in temperate climates to predict dry and wet weather, usually hanging a harvested pine cone from some string outside to measure the humidity of the air.

===Araucariaceae===
[[File:Pinhas.jpg|thumb|''[[Araucaria angustifolia]]'' cones and nuts]]
Members of the [[Araucariaceae]] (''[[Araucaria]]'', ''[[Agathis]]'', ''[[Wollemia]]'') have the bract and seed scales fully fused and have only one ovule on each scale. The cones are spherical or nearly so, 5–30&nbsp;cm diameter, and mature in 18 months. For most species they disintegrate at maturity to release the seeds, although in some such as ''[[Araucaria bidwillii]]'', the cone weighing up to {{convert|10|kg|lb}} is shed intact. In ''Agathis'', the seeds are winged and separate readily from the seed scale, but in the other two genera, the seed is wingless and fused to the scale.

===Podocarpaceae===
[[Image:Podocarpus macrophyllus (seed s3).jpg|left|thumb|Berry-like Podocarpus cone]]
The cones of the [[Podocarpaceae]] are similar in function, though not in development, to those of the Taxaceae (q.v. below), being berry-like with the scales highly modified, evolved to attract birds into dispersing the seeds. In most of the genera, two to ten or more scales are fused together into a usually swollen, brightly coloured, soft, edible fleshy [[aril]]. Usually, only one or two scales at the apex of the cone are fertile, each bearing a single wingless seed, but in ''[[Saxegothaea]]'' several scales may be fertile. The fleshy scale complex is 0.5–3&nbsp;cm long, and the seeds 4–10&nbsp;mm long. In some genera (e.g. ''[[Prumnopitys]]''), the scales are minute and not fleshy, but the seed coat develops a fleshy layer instead, the cone having the appearance of one to three small [[plum]]s on a central stem. The seeds have a hard coat evolved to resist digestion in the bird's stomach.
 {{clear|left}}

===Cupressaceae===
Members of the [[cupressaceae|cypress family]] ([[cupressus|cypresses]], [[thuja|arborvitae]], [[juniper]]s, [[Sequoia sempervirens|redwoods]], etc.) differ in that the bract and seed scales are fully fused, with the bract visible as no more than a small lump or spine on the scale. The botanical term [[galbulus]] (plural galbuli; from the [[Latin]] for a cypress cone) is sometimes used instead of strobilus for members of this family. The female cones have one to 20 ovules on each scale. They often have peltate scales, as opposed to the imbricate cones described above, though some have imbricate scales. The cones are usually small, {{convert|0.3|–|6|cm|in|abbr=in|disp=or|frac=32}} long, and often spherical or nearly so, like those of [[Nootka cypress]], while others, such as [[western redcedar]] and [[California incense-cedar]], are narrow. The scales are arranged either spirally, or in decussate whorls of two (opposite pairs) or three, rarely four. The genera with spiral scale arrangement were often treated in a separate family (Taxodiaceae) in the past. In most of the genera, the cones are woody and the seeds have two narrow wings (one along each side of the seed), but in three genera (''[[Platycladus]], [[Microbiota (plant)|Microbiota]]'' and ''[[Juniperus]]''), the seeds are wingless, and in ''Juniperus'', the cones are fleshy and [[Berry (botany)|berry]]-like (known as {{linktext|galbuli}}).
<gallery>
File:Sequoiadendron giganteum MHNT.BOT.2004.0.191.jpg|[[Sequoiadendron giganteum|Giant sequoia]] cones
File:Cupressus nootkatensis 43603.jpg|Spherical cone of Nootka cypress (''Cupressus nootkatensis'')
File:Calocedrus decurrens MHNT.BOT.2004.0.810.jpg|Long slender cones and winged seeds of California incense-cedar (''Calocedrus decurrens'')
File:Platycladus orientalis MHNT.BOT.2007.52.7.jpg|Cones and wingless seeds of [[Chinese arborvitae]] (''Platycladus orientalis'')
File:Juniperus_communis_cones.jpg|Berry-like cones of [[common juniper]] (''Juniperus communis'')
</gallery>

===Sciadopityaceae===
The cones and seeds of ''[[Sciadopitys]]'' (the only member of the family) are similar to those of some Cupressaceae, but larger, 6–11&nbsp;cm long; the scales are imbricate and spirally arranged, and have 5-9 ovules on each scale.

===Taxaceae and Cephalotaxaceae===
[[Image:Taxus baccata MHNT.jpg|left|thumb|Berry-like [[Taxus baccata|yew]] cone]]
Members of the [[Taxaceae|yew family]] and the closely related [[Cephalotaxaceae]] have the most highly modified cones of any conifer. There is only one scale in the female cone, with a single poisonous ovule. The scale develops into a soft, brightly coloured sweet, juicy, berry-like aril which partly encloses the deadly seed. The seed alone is poisonous. The whole 'berry' with the seed is eaten by birds, which digest the sugar-rich scale and pass the hard seed undamaged in their droppings, so dispersing the seed far from the parent plant.

=== Welwitschiaceae ===

''[[Welwitschia]]'' is unique cone-bearing plant is not considered a conifer but belongs in the order [[Welwitschiales]]. ''Welwitschia mirabilis'' is often called a [[living fossil]]<ref>Flowering Plants of Africa 57:2-8(2001)</ref> and is the only species in its genus, which is the only genus in its family, which is the only family in its order. The male cones are on male plants, and female cones on female plants. After emergence of the two [[cotyledons]], it sets only two more leaves. Those two leaves then continue to grow longer from their base, much like fingernails. This allows it great drought tolerance, which is likely why it has survived in the desert of [[Namibia]], while all other representatives from its order are extinct.<ref>A. Lewington & E. Parker (1999). ''Ancient Trees: Trees that Live for a Thousand Years''. Collins & Brown Ltd. {{ISBN|1-85585-704-9}}.</ref>

==Location and distribution==
[[File:Male cone of Cedar of Lebanon.JPG|thumb|Male cone of cedar of Lebanon]]
[[File:Larch with unusual terminal branch.JPG|thumb|left|Larch cone with unusual branch growth from tip]]
Most conifer species are [[monoecy|monoecious]], with male and female cones occur on the same plant ([[tree]] or [[shrub]]), with female usually on the higher branches towards the top of the plant. This distribution is thought to improve chances of [[cross-fertilization]], as pollen is unlikely to be blown vertically upward within the crown of one plant, but can drift slowly upward in the wind, blowing from low on one plant to higher on another plant. In some conifers, male cones additionally often grow clustered in large numbers together, while female cones are more often produced singly or in only small clusters. Some, such as ''[[Araucaria araucana]]'' and ''[[Taxus baccata]]'', are [[dioecy|dioecious]], with the male and female cones usually on separate trees, but even in normally dioecious species, scattered individuals may produce cones of both sexes, or change which sex cones they produce over time.<ref name="Rushforth"/> Some pines, notably ''[[Pinus discolor]]'', are subdioecious, with individuals producing cones of both sexes, but with each tree producing either predominantly male, or predominantly female, cones, and only a few of the other sex.<ref>{{cite journal |last1=McCormick |first1=Jack |last2=Andresen |first2=John W. |title=A Subdioecioius Population of Pinus cembroides in Southeast Arizona |journal=Ohio Journal of Science |date=1963 |volume=63 |issue=4 |pages=159–163}}</ref><ref>{{cite journal | last1=Bailey | first1=D. K. | last2=Hawksworth | first2=F. W. | title=Pinyons of the Chihuahuan Desert Region |journal=Phytologia | volume=44 |pages=129–133 | date=1979 | issn=0031-9430 | url=https://www.biodiversitylibrary.org/page/13044791 | access-date=2025-02-12}}</ref>

A characteristic arrangement of pines is that the male cones are located at the base of the branch, while the female at the tip (of the same or a different branchlet). However, in larches and cedars, both types of cones are always at the tips of short shoots, while both sexes of fir cones are always from side buds, never terminal. There is also some diversity in bearing in Cupressaceae. Some ''[[Cupressus]]'' for instance, have little or no differentiation in the positions of male and female cones.
{{Clear}}

===Crop potential===
Cone crop potential can be predicted in various ways. An early indication of a potential crop can be a period of abnormally hot, dry weather at the time of bud differentiation, particularly if the current and preceding cone crops have been poor (Nienstaedt and Zasada 1990).<ref name="nien5">{{Silvics |last1=Nienstaedt |first1=Hans |first2=John C. |last2=Zasada |volume=1 |genus=Picea |species=glauca}}</ref> Estimates of cone crop potential can be made by counting female reproductive buds in fall or winter, and an experienced observer can detect the subtle morphological differences and distinguish between reproductive buds and vegetative buds (Eis 1967b).<ref name="eis2">Eis, S. 1967b. Cone crops of white and black spruce are predictable. For. Chron. 43(3):247–252.</ref>

White spruce seed collection is expensive, and collection from cone caches of [[red squirrel]]s is probably the cheapest method. The viability of seed from cached cones does not vary during current caching, but viability drops essentially to zero after being in caches for 1 or 2 years.<ref name="wagg">Wagg, J.W.B. 1964. Viability of white spruce seed from squirrel-cut cones. For. Chron. 40(1):98–110.</ref> Collection of cones in seed orchards has been facilitated by the counter-intuitive technique of "topping" and collection of cones from the severed crown tops at one-third the cost of collection from untopped trees and without decreasing cone production.<ref name="slay">Slayton, S.H. 1969. A new technique for cone collection. USDA, For. Serv., Tree Plant. Notes 20(3):13. (Cited in Nienstaedt and Zasada 1990).</ref><ref name="nien1">Nienstaedt, H. 1981. Top pruning white spruce seed orchard grafts does not reduce cone production. USDA, For. Serv., Tree Plant. Notes 32(2):9–13. (Cited in Coates et al. 1994).</ref>

==Pseudocones==
<gallery>
Developing Pineapple Gall.JPG|Developing pineapple pseudocone galls on Norway spruce
Sitka Spruce with Pineapple Gall.JPG|Pseudocone on Sitka spruce
Spruce gall on Picea breweriana. - Flickr - theforestprimeval.jpg|Pseudocone on Brewer's spruce
</gallery>
Most species of spruce are prone to the formation of [[Adelges|pineapple gall]] pseudocones caused by the woolly aphids in the genus ''[[Adelges]]''. These are not cones, although they closely resemble them. [[Alder]] (''Alnus'') trees are not conifers, but their mature seed bearing catkins closely resemble cones.{{Clear}}

{{Anchor|Cone cows}}

==Cultural uses==
[[Image:Blason ville fr Sarrant (Gers).svg|thumb|Cone in the coat of arms of [[Sarrant]].]]
Because of their widespread occurrence, conifer cones have been a traditional part of the [[handicraft|arts and crafts]] of cultures where conifers are common. Examples of their use includes seasonal wreaths and decorations, fire starters, bird feeders, toys, etc.<ref>{{cite web |url=http://familycrafts.about.com/od/pineconecrafts/Pine_Cone_Craft_Projects.htm |title=Pine Cone Craft Projects |publisher=Familycrafts.about.com |date=2013-08-23 |access-date=2013-08-30 |archive-date=2013-05-11 |archive-url=https://web.archive.org/web/20130511004635/http://familycrafts.about.com/od/pineconecrafts/Pine_Cone_Craft_Projects.htm |url-status=dead }}</ref> An intriguing derivation of the [[impossible bottle]] [[mechanical puzzle]] takes advantage of the fact that pine cones open and close based on their level of dryness.  In constructing a display, a closed, damp cone of suitable size is inserted into a narrow-mouthed bottle and allowed to open upon drying.<ref>{{cite web|author=zupperzipper |url=http://www.instructables.com/id/pine-cones-in-a-bottle |title=Pine Cone in the Bottle Display |publisher=Instructables |date=2008-07-13 |access-date=2013-08-30}}</ref>

Cone cows are traditional homemade toys, made by children using material found in nature. The most common design is a spruce or pine cone with sticks or [[match]]es for legs, which can easily be attached by forcing them between the cone scales. Playing with cone cows often includes building an animal enclosure from sticks. For the most part, cone cows have been displaced by manufactured toys, at least in affluent countries, but the creation of cone cows still enjoys some popularity as an outdoor activity for children. Cone cows are a part of children's culture in [[Finland]] where they are known as ''käpylehmä'' and [[Sweden]] where they are known as ''kottkor'' (cone animals).

In Sweden, a video game was released in which the player may build virtual cone cows.<ref>[http://www.aftonbladet.se/nojesbladet/recensioner/spel/article109235.ab Kapsyljakt med Anki och Pytte review] "You build cone cows"</ref> Swedish artist [[Lasse Åberg]] has created artwork with cone cows, which has been included in an [[alphabet book]]<ref>[http://www.panorstedt.se/templates/raben/Book.aspx?id=23250 Åbergs ABC]{{dead link|date=August 2017 |bot=InternetArchiveBot |fix-attempted=yes }} "a little book about cone cows, small fears, swarming helicopters and such"</ref> and featured on a Swedish [[postage stamp]] among other classic toys.<ref>[http://cws.huginonline.com/P/134112/PR/200010/886184_2_12.html Posten.se press release] {{webarchive|url=https://web.archive.org/web/20100913223636/http://cws.huginonline.com/P/134112/PR/200010/886184_2_12.html |date=2010-09-13 }} "Motives include [..] life in the countryside, which shows both the cone cow, a doll in a traditional outfit, and a horse and carriage toy"</ref><ref>[http://www.postmuseum.posten.se/frimbas/frimarks/F2000043.asp Livet på landet] image of the stamp from the official postal site</ref>[[Image:Cortile della Pigna pine cone 2.jpg|thumb|Pine cone statue from the Cortile della Pigna of the [[Vatican Museums]]]]
[[Image:Koper fountain.JPG|thumb|[[Da Ponte fountain]] in [[Koper]]]]
Cones are used as decorative elements in architecture such as on top of the posts surrounding [[Koper]]'s [[Da Ponte Fountain]], the central element of the {{Lang|it|[[Fontana della Pigna]]|italic=no}} in [[Rome]], or a bronze cone in the narthex of the [[Aachen Cathedral]]. Cones are occasionally used as a charge in [[heraldry|heraldic]] coats of arms. In some parts of [[Russia]] and [[Georgia (country)|Georgia]], immature pine cones are harvested in late spring and boiled to make sweet [[Fruit preserves|preserves]].<ref name="atlasobscura">{{Cite web|url=https://www.atlasobscura.com/foods/pine-cone-preserves|title=Pine Cone Preserves|website=Atlas Obscura}}</ref>[[File:Armes de la famille Parthon de Von.svg|thumb|[[Coat of arms]] of the [[Parthon de Von family]] featuring three pine cones]]
The [[pineal gland]] is named after the pine cone.<ref name="oed">{{cite web |title=Pineal (as an adjective) |url=https://www.etymonline.com/word/pineal |publisher=Online Etymology Dictionary, Douglas Harper |access-date=27 October 2018 |date=2018}}</ref> Pine cones were also used as symbols of fertility in ancient Assyrian art. In Christian symbolism, they are closely related to the [[Tree of life#Christianity|tree of life]].<ref>The Continuum Encyclopedia of Symbols by Udo Becker (2000) Page 234</ref>

==Gallery==
<gallery class="center" perrow="6" mode="packed">
Image:Cedar of Lebanon cone.JPG|Female cone of a [[Cedrus libani|Lebanese cedar]]
Image:Pine_cones,_immature_female.jpg|Immature female pine cone
Image:Japanese Larch pollen cone, Cardiff, Wales.jpg|Pollen cone of a [[Japanese larch]]
Image:Pineapple gall.JPG|[[Pineapple gall]] on [[Sitka spruce]] caused by Adelges abietis.
Image:pine_cones,_male_and_female.jpg|Dozens of male cones (orange and flower-like) occur in a cluster; the female cone is still immature (olive green). [[Lodgepole pine]].
Image:loblolly_pine_strobili.jpg|Young female cones of [[loblolly pine]] receptive for pollination.
File:Loblolly male flowers.jpg|[[Loblolly pine]] male cones ready to cast pollen.
Image:shortleaf_cone.jpg|Cross section of maturing [[shortleaf pine]] cone showing seeds (arrows).
Image:ages_of_pine_cones.jpg|[[Loblolly pine]] branch with cones of different ages; two-year old cones will disperse seeds during fall and winter.
Image:Pinus canariensis (male) in Presa de las Niñas 02.jpg|[[Pinus canariensis]] male cone in [[Gran Canaria]].
File:Abies bracteata 01 Cone Peak.jpg|The cone structure of [[Abies bracteata]]
File:Top of a pine cone.jpg|Top of a pine cone
</gallery>

== Visualisation of a ''Cupressus'' cone by micro computed tomography ==
<gallery>
File:Cupressus strobilus ("cone"), front and back.jpg|Photographs of the cone, front and back
File:ΜCT Cupressus strobilus 01.jpg|3D visualisation of the cone
File:ΜCT Cupressus strobilus 02 - 3D section series.jpg|Various sections through the 3D object
File:ΜCT Cupressus strobilus 03 semi-transparent.jpg|Semi-transparent view
File:ΜCT Cupressus strobilus 04 mid section.jpg|Midsection
File:ΜCT of a cupressus strobilus, spiral 3D flight.ogg|Flight around the 3D object
File:ΜCT of a cupressus strobilus, frontal flight through image stack.ogg|Flight through the μCT image stack, lateral view
File:ΜCT of a cupressus strobilus, top flight through image stack.ogg|Flight through the μCT image stack, top view
</gallery>

==See also==
* [[Conifer nuts]]
* [[Parastichy]]

==References==
{{Reflist}}

==External links==
{{Commons category|Conifer cones}}
*[http://www.pinetum.org/cones/mpfcones.htm Arboretum de Villardebelle] Images of various conifer-cones
*[https://web.archive.org/web/20060425221129/http://www.cas.vanderbilt.edu/bioimages/pages/cones.htm Gymnosperm cone images at bioimages.vanderbilt.edu]

{{Authority control}}

{{DEFAULTSORT:Conifer Cone}}
[[Category:Conifers]]
[[Category:Plant morphology]]
[[Category:Articles containing video clips]]
[[Category:Heraldic charges]]